Anais da Academia Brasileira de Ciências ISSN: 0001-3765 [email protected] Academia Brasileira de Ciências Brasil

MEDINA, MARIA CAMILA; HOYOS-GÓMEZ, SAÚL E.; DEMARCO, DIEGO; TUBERQUIA, DINO Peculiar anatomical traits, high durability, and potential ornamental use of as fresh foliage Anais da Academia Brasileira de Ciências, vol. 89, núm. 3, 2017, pp. 2399-2410 Academia Brasileira de Ciências Rio de Janeiro, Brasil

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How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative Anais da Academia Brasileira de Ciências (2017) 89(3 Suppl.): 2399-2410 (Annals of the Brazilian Academy of Sciences) Printed version ISSN 0001-3765 / Online version ISSN 1678-2690 http://dx.doi.org/10.1590/0001-376520172-170128 www.scielo.br/aabc | www.fb.com/aabcjournal

Peculiar anatomical traits, high durability, and potential ornamental use of Cyclanthaceae as fresh foliage

MARIA CAMILA MEDINA1,3, SAÚL E. HOYOS-GÓMEZ2, DIEGO DEMARCO1,3 and DINO TUBERQUIA1

1Programa de Biología, Universidad CES, Medellín, Calle 10 A No. 22-4, Antioquia, Colombia 2Corporación Fragmento, Medellín, Cra 77 No 33 A 61, Antioquia, Colombia 3Departamento de Botânica, Instituto de Biociências, Universidade de São Paulo, Caixa Postal 11461, 05508-090 São Paulo, SP, Brazil

Manuscript received on February 22, 2017; accepted for publication on July 3, 2017

ABSTRACT Wild species with potential for use as ornamental green foliage are often reported. Nevertheless, there are few studies evaluating morphological and anatomical aspects associated with this use. Preliminary trials suggest that the species of Sphaeradenia (Sander ex Mast.) Harling, a Cyclantaceae that occurs from Costa Rica to Peru, is suitable to be used as fresh foliage because of attractiveness of their leaves in vase and its durability without showing tissue damage, compared with other genera of this family. Through the study of anatomical characters, it is possible to predict the potential of a species that can serve as ornamental foliage. In this study, three foliar anatomical features (thickness of the cuticle and leaf, and percentage of fibers in the mesophyll) of seven Cyclanthaceae species were evaluated, using anatomical techniques and image analysis with the LUCIA® and ImageJ softwares. Statistical analysis of the results suggests that this high durability is associated with the thickness of the cuticle and mesophyll, but not with the amount of fibers in the leaf. Key words: anatomy, cuticle, fibers, mesophyll, Sphaeradenia laucheana.

INTRODUCTION whose juvenile forms have been widely used (Chen et al. 2002a, b). Currently, more than 500 species with various In general, foliage are resistant plants shapes, colors, textures and styles, mostly from tropical and subtropical environments, form that grows in greenhouses shadows. Their leaves the basis of the industry of foliage plants. Some are quite attractive; as a result, they are used of them are found in desert environments as mainly as live specimens for interior decoration, succulent plants and cactus. They are rarely found rather than perishable or fruits. They grow in temperate climates. The most significant native under the canopy of trees on the forest floor or live foliage plant in temperate climates is Hedera helix, on trees as vines or epiphytes. Their tolerances to

Correspondence to: María Camila Medina light are low, and are sensitive to high temperatures E-mail: [email protected] (Chen et al. 2002b, 2005, 2010).

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The basic function of the ornamental Kingdom who account for about 50% of spending horticulture lies in the satisfying of the aesthetic on ornamental (Díaz M et al. 2002, Díaz M. 2006). needs of man. Today it is considered one of the Markets for foliage plants require quality most attractive businesses because it can provide standards for export. Therefore these plants must high income per unit area. Beautifying indoor meet conditions to be used in the production environments and indoor air purification have of bouquets and other floral arrangements, for become key elements for the advancement of example, the time it can be kept in good condition the use foliage plants, because as it has been after being harvested (Yuste Molina 1992, Gutiérrez demonstrated that they remove pollutants such et al. 2007). One of the species for export Rumohra as formaldehyde, benzene, toluene, xylene, and adiantiformis, commonly known as “leather fern”, others from air (Wolverton et al. 1989, Mosaddegh that maintains its quality and freshness for a long et al. 2014)which are probably more polluted than period of time (25 days) after it has been harvested outside the buildings. High levels of volatile organic and packaged (López Marin et al. 2006). compounds (VOCs. Another important factor in On the other hand, the non-forestry use of forests the steady growth of the industry foliage plants is an alternative to the problem of deforestation (López Marin et al. 2006 ), particularly in regions is the continuous adjunction of new plants and close to urban centers (García and Polaina 2010). the development of new cultivars, which further Cyclanthaceae are almost exclusively confined expand the options for using of foliage plants, like to the Neotropical parts of the world. Nearly all the houseplants (Chen et al. 2002b). Cyclanthaceae require wet and more or less shady Over the years, some species belonging to habitats. They are therefore mainly found in the several genera have been added to the list of species rain forests, both in the tropical rain forests and used in industry of foliage plants. Aglaonema, in the montane rain forest. Just like the leaves of Dieffenbachia, Dracaena, Epipremnum, Ficus, many and Araceae, the juvenile leaves Hedera, Peperomia, Philodendron, Sansevieria, of the Cyclanthaceae come very close to the Syngonium, and plants of several genera of the ancestral leaf type of the . The leaf family Arecaceae, Polypodiaceae, Poaceae, blades vary greatly in respect to their size, shape, Cyperaceae, among others, have been the backbone venation, consistence and color. They are also of the industry for at least half a century. Only generally more or less deeply cleft, or divided by the Araceae family includes at least 10 important three rather deep incision into four wedge-shaped genera of ornamentals foliage plants (Henny et al. lobes, which in their turn are regularly dentate or 2008). lobate in their upper part, or can be entire (Harling Worldwide production of plants and flowers 1958, Wilder 1981). The mesophyll is sometimes has been extending over the years. Numerous distinctly differentiated into palisade and spongy production centers are localized in developing parenchyma but in many species this differentiation countries and they supply, on a regular basis to is lacking (Wilder 1985a). the principal consumers such as Japan, the United A comparative study on S. laucheana and six Stated and Western Europe. These countries other Cyclanthaceae species with different ranges absorb about 75% of the worldwide production of geographical distribution, some of them already of ornamentals, worth of 80.000 million euros. used as ornamentals foliage and landscaping, was About 36.000 millions of euros correspond to conducted. The evaluation is to help determine the Europe, being Germany, France and the United anatomical characteristics that would let explain

An Acad Bras Cienc (2017) 89 (3 Suppl.) ANATOMICAL TRAITS OF ORNAMENTAL CYCLANTHACEAE 2401 its durability and potential use as foliage, thus TABLE I proposing a study mechanism for prospecting new Coding of the Cyclanthaceae specimens studied. Collection Statistical Species species with potential use as foliage. Number Coding Two sets of variables were taken into account; DT-3415 Asplundia sp. Acalle 1) durability, determined by the lifetime or duration DT-3413 Asplundia flavovaginata Aflavo in vase, also known as “vase life method” described AC-12800 drudei Cdrudei by Gutiérrez et al. (2007); 2) relevant anatomical AC-12801 Carludovica palmata Cpalma features, namely thickness of the cuticle and AC-12800 Dicranopygium calimense Dcalim mesophyll thickness and the percentage of fibers DT-3611 Sphaeradenia danielii Sdaniel present in the mesophyll and their possible impact DT-3612 Sphaeradenia cuatrecasana Scuatrec on the vase durability. DT-3610 Sphaeradenia laucheana Slauch

MATERIALS AND METHODS cuticle and mesophyll thickness and the number of fibers, fiber strand number and percentage of SPECIES AND STUDY AREA fibers in the mesophyll; of which, measurements of The studied species were: Asplundia sp. Tuberquia, length and area were performed. Asplundia flavovaginata Harling, Carludovica The sections were examined with an optical drudei Mast, Carludovica palmata Ruiz & Pav, microscope Nikon Eclipse TE2000 with 100X Dicranopygium calimense Harling, Sphaeradenia to 400X magnification. Digital images, three for danielii Harling, Sphaeradenia cuatrecasana each leaf area were captured using a Nikon camera Harling and Sphaeradenia laucheana (Sander ex Digital Sight (DS-5M). Each digital image was Mast.) Harling. taken from a different anatomical cut. The study area is located in the Valle de The measurements were made with Aburrá, Antioquia department. S. laucheana, S. image analysis software LUCIA® Version 5.3 cuatrecasana and S. danielii samples for anatomical (Laboratory Imaging, Prague, Czech Republic) and analysis and durability tests proposed for this study they were: 15 cuticle length measurements for each were collected from a sample of relict forest in the and 15 mesophyll thickness, measurements for Astilleros plantation terrain, San Antonio de Prado, each individual, and two measurements of the area Medellin. D. calimense, A. sp., A. flavovaginata, occupied by the fibers in the mesophyll; this area C. drudei and C. palmata samples were collected was measured delimiting the outside of the band from a living collection of Cyclanthaceae in the with a freehand line using the LUCIA® software, Jardín Botánico Joaquín Antonio Uribe, Medellin. which yields the value of the delineated area. The number of fibers strands, number of fibers and the SECTIONING AND ANATOMICAL MEASUREMENTS area occupied by the mesophyll in a visual field An identification code was assigned to each species was registered.

(Table I). The samples were preserved in 90% PLATE MOUNTING alcohol and taken to the Laboratory of Histology of the Universidad CES for further processing. Given that most of the species considered in this In order to conduct the anatomical study have leaves longer than 1 m, the leaf was characterization, free-hand cross-sections were divided into three areas: apical, middle and basal, made in order to compare the anatomical features and measurements were done on cuticle, mesophyll supposedly related to durability in vase, namely and fibers. For each zone of the leaf, histological

An Acad Bras Cienc (2017) 89 (3 Suppl.) 2402 MARIA CAMILA MEDINA et al. sections were performed in triplicate. The leaf value is multiplied by one hundred and divided cross sections were done freehand fixed with CaCl2 by the mesophyll total area of same cross section. 16% (Keating 2014) and stained with Cresyl Violet Additionally it was done counting the number of Acetate (CVA). The blades were mounted semi- fibers and the number of strands for each cross- permanent. section fibers

DURABILITY TEST STATISTICAL ANALYSIS

The durability test was based on the proposed The thickness of the cuticle (cut), mesophyll criteria by Gutiérrez et al. (2007): a specimen is thickness (mes) and percentage of fibers (pfiber), reputed to have good performance of vase life between species and between areas of the leaf if the material retains its initial characteristics blade were processed using the free software (“R after 15 days of evaluation period; i.e. there is no development Core Team” 2011). collapse of stem, dehydration, damage by fungi, Each variable was analyzed using a two discoloration, geotropism or necrosis. For this way ANOVA including species factor and the test, two leafs, including 0.15 m of petiole, from area factor of the sheet. The model includes the different individuals of each species, were placed interaction between the two factors. The Bonferroni in containers with water and subjected to daily adjustment was used for multiple comparisons monitoring; the external appearance of the leaf between pairs of species. The ANOVA model was recorded in photographs every 10 days (Fig. assumptions are validated by the residual graph 1). Considering that for ornamental foliage can technique. The cuticle (cut) and mesophyll (mes) be used, it must have duration of two weeks, the values were transformed to natural logarithm in durability test conducted for 5 weeks for detailed order to correct for heterogeneity of variances. The observation of each species. resulting averages for these variables (cut and mes) were transformed back to its original scale. DURABILITY TEST VERSUS CUTICLE AND MESOPHYLL THICKNESS, AND PERCENTAGE OF RESULTS MESOPHYLL FIBERS DURABILITY TEST Three anatomical variables were selected would be related to high durability in vase of each species Two leaves from different individuals of the same (cuticle and mesophyll thickness, and mesophyll species were subjected to the test. The eight species percentage of fibers). To establish this association, tested showed different lengths of durability in the samples used for anatomical measurements vase, as illustrated in Fig. 1. The test time takes into and durability tests were obtained from the same account that Sphaeradenia laucheana lasts about individual. The leaf blade was divided into three three months without any harm, while other species sections: apical, middle and basal, for performing last a shorter time. It was apparent that both leaves, measurements of anatomical variables. With of the same species, showed signs of deterioration the average values of the measurements were simultaneously, during one-day interval, except performed by lineal regression between the results C. palmata whose leaves deteriorated seven days of durability results and anatomical measurements. apart (18 vs. 25 days). The main cause of removing The percentage of each cut fibers was obtained species was necrosis in the apical area of the leaves; by adding the area measurements strands fibers dehydration and leaf crinkling eliminated only S. present in the mesophyll of each cross-section, this cuatrecasana.

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(Figs. 5, 6, 7), being more evident for the species S. laucheana, which lasted 37 days and presented the highest values of cuticle thickness.

MESOPHYLL THICKNESS MEASUREMENTS

A marked difference in the thickness of the mesophyll is observed among the analyzed species (P <0.0001). S. danielii and S. laucheana were the species that showed thicker mesophylls and C. drudei showed the lowest thickness of the mesophyll (Table II) values. Mesophyll thicknesses between different areas of the leaf (apical, middle and basal) of all individuals from the eight species studied were compared. A significant difference in mesophyll thickness between different areas of the leaf, with values ranging between 200 and 600 micrometers in S. danielii and S. laucheana was observed, the Figure 1 - Durability test (vase life) of eight species of basal area being thicker in both species. Cyclanthaceae during 37 days. a. State of the leaves on the 16th day of monitoring. b. Duration times for each species. DURABILITY TEST VS. MESOPHYLL THICKNESS MEASUREMENTS CUTICLE MEASUREMENTS There is an apparent association between the mean No significant differences in the thickness of the values of mesophyll thickness and durability in vase cuticle in relation to the three areas of the leaf within (Fig. 3). The mesophyll of A. sp., S. cuatrecasana the same species; nevertheless there are differences and S. laucheana showed similar ratios with respect in the thickness of the cuticle (P <0.0001) between to the thickness of the mesophyll, but S. laucheana the eight species as shown in Table II. durability surpassed the other species. D. calimense presented the lowest values of PERCENTAGE OF FIBERS cuticle thickness. The species belonging to the genus Sphaeradenia showed the highest values The values of fiber percentage from the eight species in the cuticle thickness measurements and S. vary significantly (P = 0.0012). The species with the laucheana showed the highest values for the three higher percentage of fibers was C. palmata, with areas of the leaf (Table II). the highest value for the apical area and the lowest for the basal area. (Apical: 8.18%, Central: 7.35%, DURABILITY TEST AND THICKNESS MEASUREMENTS CUTICLE basal: 6.41%). This species had the highest number of fibers in the central region (38.75 fibers), and the Long durability (more than 15 days of vase life), largest number of bands for the three areas of the as shown by linear regressions (Fig. 2), may be sheet (apical: 13.00, central: 17.25, basal: 13.25). associated with a thicker cuticle in some species Sphaeradenia laucheana had the highest (S. cuatrecasana, S. danielii and S. laucheana,) number of fibers in the apical and basal region of

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TABLE II Mean values (micrometers) of measurements of thickness of the cuticle, mesophyll thickness, percentage and number of fibers in the mesophyll with their respective standard deviations. Species Apical ED Middle ED Basal ED A. sp. 3.59 1.10 3.89 1.08 3.87 1.08

A. flavovaginata 2.96 1.08 2.90 1.08 3.12 1.08

C. drudei 4.43 1.10 4.02 1.08 4.22 1.08

C. palmata 4.12 1.10 4.74 1.10 4.84 1.10 Cuticle D. calimense 1.87 1.08 2.17 1.08 3.22 1.08

S. cuatrecasana 5.44 1.08 8.09 1.10 6.09 1.08

S. danielii 5.24 1.08 6.46 1.08 8.02 1.08

S. laucheana 9.95 1.08 11.46 1.08 12.70 1.08

A. sp. 244.52 1.05 315.54 1.05 428.65 1.05

A. flavovaginata 191.35 1.05 246.92 1.05 335.44 1.05

C. drudei 138.77 1.05 179.07 1.05 243.27 1.05

C. palmata 145.73 1.05 188.06 1.06 255.48 1.06 Mesophyll D. calimense 160.00 1.05 206.74 1.05 280.49 1.05

S. cuatrecasana 198.25 1.05 255.83 1.05 347.54 1.05

S. danielii 256.28 1.05 330.72 1.05 449.27 1.05

S. laucheana 261.61 1.05 337.59 1.05 458.61 1.05

A. sp. 1.96 0.95 2.84 0.95 2.57 0.95

A. flavovaginata 3.76 0.95 4.64 0.95 5.63 0.95

C. drudei 4.36 0.95 5.45 0.95 4.91 0.95

C. palmata 8.18 0.95 7.35 0.95 6.41 0.95 Fiber Percentage D. calimense 3.82 0.95 3.30 0.95 4.16 0.95

S. cuatrecasana 5.78 0.95 4.95 0.95 6.66 0.95

S. danielii 2.16 0.95 2.41 0.95 7.41 0.95

S. laucheana 6.55 0.95 4.75 0.95 4.74 0.95

A. sp. 8.25 4.92 11.25 4.92 11.50 4.92

A. flavovaginata 16.50 4.92 35.00 4.92 51.25 4.92

C. drudei 18.00 4.92 29.50 4.92 33.25 4.92

C. palmata 35.00 4.92 38.75 4.92 35.50 4.92 Number of fibers D. calimense 13.50 4.92 12.08 4.92 18.75 4.92

S. danielii 17.00 4.92 20.50 4.92 63.00 4.92

S. laucheana 47.50 4.92 34.25 4.92 65.25 4.92

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TABLE II (continuation) Species Apical ED Middle ED Basal ED A. sp. 5.00 1.43 5.25 1.43 4.50 1.43 A. flavovaginata 6.50 1.43 8.50 1.43 11.00 1.43 C. drudei 6.50 1.43 14.50 1.43 14.00 1.43 Number of bands C. palmata 13.00 1.43 17.25 1.43 13.25 1.43 D. calimense 4.75 1.43 4.00 1.43 5.50 1.43 S. danielii 4.00 1.43 6.00 1.43 12.50 1.43 S. laucheana 5.00 1.43 4.50 1.43 9.50 1.43

Figure 2 - Relationship between the mean values of cuticle thickness and durability in vase for eight species of Cyclanthaceae. See name species in Table I. the leaf (apical: 47.50, basal: 65.25), outnumbering having a shape, amount and distribution of fibers C. palmata (apical: 35.00, basal: 35.50). It should obviously different (Figs. 9, 10). Some species have be realized that the bands of fibers vary in sizes and the highest percentage of fibers in the apical region shapes, depending on the morphology of the fibers (C. palmata and S. laucheana), other ones in the comprising it. In S. laucheana, few bands with central region (A. sp. and C. drudei), and other ones numerous fibers of small size and smaller lumens in the basal region (S. cuatrecasana, S. danielii, A. are observed (Fig. 7). In C. palmata, numerous flavovaginata and D. calimense) as per Table II. bands with few large fibers and large lumens On the other hand, some species have scarce fibers are observed (Fig. 9), and also present a higher bands in mesophyll (Fig. 8). percentage of fibers and number of bands, and S. DURABILITY TEST VS. PERCENTAGE OF FIBERS laucheana a larger number of fibers. The percentage and organization of the There is not a high correlation between the durability fibers did not present a uniform distribution in test and the percentage of fibers, as shown in Figure the mesophyll, since these characteristics varied 4. Some species have a higher percentage of fibers according to each species. This is clearly observed in the apical area decreasing towards the basal zone in the two studied species of the genus Carludovica, and in other ones the opposite case occurs.

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Figure 3 - Relationship between the mean values of mesophyll thickness and durability in vase for eight species of Cyclanthaceae. See names of each species in Table I.

Figure 4 - Mean values of the percentage of fiber compared with durability in vase for eight species of Cyclanthaceae. See species names in Table I.

Carludovica palmata presents the highest because of its high durability in vase. The results of percentage of fibers in the mesophyll in the basal this study suggest that high durability is associated and central areas of the leaf, reaching a 20-day with the thickness of the cuticle and mesophyll average durability, but A. sp. and S. cuatrecasana compared to other species of the Cyclanthaceae present the same durability; this later one with family and to a lesser extent this durability is the lowest percentage of fibers, and S. laucheana associated with the percentage of sclerenchyma presents a medium percentage of fibers, and the fibers. longest durability. In Cyclanthaceae family, a combination of layers appears in the cuticle, which causes DISCUSSION it to observe much thicker (Wilder 1985a). This Sphaeradenia laucheana is a species with potential combination occurs as two discrete regions or for use as ornamental foliage (Gutiérrez et al. 2007) fields: the first, called no cutinized region and the

An Acad Bras Cienc (2017) 89 (3 Suppl.) ANATOMICAL TRAITS OF ORNAMENTAL CYCLANTHACEAE 2407 second the cutinized region, both covered by a storing water. This allows them to have greater cuticle sensu stricto. The cuticle can be thicker in desiccation tolerance. Wilder (1985c) says that in the abaxial or adaxial region or may be equal in all species the mesophyll contains strands of one both epidermises. The combined thickness of these or more fibers, which normally are parallel to the layers varies depending on the species. Harling longitudinal veins. (1958) reports that the leaves in the species of the In all species studied, the fibers appear group he called informally Sphaeradenia (with organized in strands, scattered throughout all three genera) are more coriaceous than those of the regions of the mesophyll (adaxial, central and group called Asplundia (six genera). Anatomical abaxial). Fiber strands occur in larger quantity studies by Wilder (1985a) show that some species in the adaxial region in S. cuatrecasana and, S. of the genera Dicranopygium and Asplundia have danielii. The fiber strands occur in similar amounts the lowest thickness of cuticle and most species in both regions of mesophyll in D. calimense, A. sp., of the genus Sphaeradenia present the largest and A. flavovaginata, in the apical area of the leaf. thickness. These results agree with those obtained Fiber strands were observed scattered throughout in this study, where three species of the genus the mesophyll in C. drudei and C. palmata as they Sphaeradenia showed the highest values of cuticle were observed by Wilder (1985c), who reported thickness as shown in Table II. the fiber strands concentrated mainly in the abaxial It is known that most of species of Sphaeradenia, region of the leaf blade, in some species of this grow above 2000m altitude, and S. laucheana family. reaches extreme altitudinal limits, even up to 3700 It should be considered that the sclerenchyma m altitude (Eriksson 1995). Thick cuticles in some cell morphology varies in shape and size according plants could be related to ecological requirements to the species (Wilder 1985c). In some species the in high mountain environments (Hlwatika and Bhat fibers had wide lumens that occupy a large area 2002). of the cell, and thus occupying large areas in the Some species of Cyclanthaceae, especially mesophyll (C. palmata, A. sp. and D. calimense). Sphaeradenia laucheana, show features that allow Therefore, when measuring areas occupied by them to be resistant to desiccation, which enhance fibers in these three species area, large areas were the possibility of being used as ornamentals. obtained. Fibers with medium size lumens and Their features include specialized cells as fibers, generally small for these two species were observed cells containing crystals (this occur solely in the in A. flavovaginata, C. drudei and S. laucheana. The mesophyll) (Wilder 1985b), tanniniferous and strands of these fibers also have medium sizes, from internal secretory tissues as mucilage canals 2 to 8 fibers per strand. In other cases, as occur in or cavities are found in all the species of family species of the genus Sphaeradenia; fibers presented taking place in the stem, petiole, the peduncle, lumens of small sizes, but with more lignified inflorescence, often also in the root and in the area. Wilder (1985c) reported that depending on costa of the leaf blade (Harling 1958). According the species, mesophyll fibers might generally have to (Fahn and Cutler 1992) the mucilage in plants is narrow or broad lumens. In some species, some suggested to serve for water retention, because of fibers of larger veins are broader and exhibit wider their great ability to imbibe water, it was assumed lumens than those of the mesophyll. These fibers that the mucilage plays a role in regulation of the were grouped in large strands, up to 20 fibers per water economy in plants. It has been suggested band; these strands occur in small quantities in the that the presence of fibers may have a role in mesophyll; when the areas occupied by the fibers in

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Figures 5-10 - Cross-sections of leaves of Cyclanthaceae. 5. S. cuatrecasana cuticle 100x. 6. S. danielli cuticle 100x 7. S. laucheana fiber strands and cuticle 40x.100x 8. A. flavovaginata fibers strands 9. C. palmata fiber strands 40x. 10. C. drudeii fiber strands 40x.

An Acad Bras Cienc (2017) 89 (3 Suppl.) ANATOMICAL TRAITS OF ORNAMENTAL CYCLANTHACEAE 2409 the mesophyll are compared between species, those ACKNOWLEDGMENTS of the genus Sphaeradenia, show the lowest ones. We want to express our appreciation to the Botanical The percentage of fibers present in the Garden Joaquín Antonio Uribe (Medellín) for the mesophyll is not, apparently, a determining factor in access granted to live specimens; Universidad CES the high or low vase durability. C. palmata presents for the logistical support; Yeison Montoya for his the highest value in percentage of fibers but a help in the field; the Biological Engineering Magda medium vase life; S. laucheana presents a medium Ledesma Eraso, for advice in managing LUCIA® fiber percentage but the maximum vase life. A. Software, and Biologist Pablo Andres Guzmán for sp. presents the minimum fiber percentage but its advice on statistical analysis. vase life is comparable with that of C. palmata. These observations highlight the importance of REFERENCES measurements to be performed accompanied CHEN J ET AL. 2002a. Production and Interior Performances by close observation of the organization and of Tropical Ornamental Foliage Plants Grown in Container morphology of the fibers in the mesophyll. Substrates Amended with Composts. Compost Sci Util 10: 217-225. This study does not support the conclusion CHEN J ET AL. 2002b. Development of New Foliage Plant that the durability is directly related to any of Cultivars. Trends in new crops and new uses, p. 466-472. the anatomical attributes evaluated in particular. CHEN J ET AL. 2005. Response of Tropical Foliage Plants to Interior Low Light Conditions. Acta Hort, p. 51-56. It may suggest that the high vase durability CHEN J ET AL. 2010. The Foliage Plant Industry. of Sphaeradenia laucheana has to do with an DÍAZ MJA. 2006. Diagnóstico de la cadena productiva de interaction of different variables; cuticle thickness, heliconias y follajes en los departamentos del eje cafetero y Valle del Cauca ( Colombia ) Consultor Biocomercio- mesophyll thickness and percentage of fibers BTFP, p. 1-46. present in the mesophyll. Some species included DÍAZ MJA ET AL. 2002. Sondeo del mercado internacional in this study, grow in low-middle elevations (C. de Heliconias y Follajes Tropicales. ERIKSSON R. 1995. The genus Sphaeradenia (Cyclanthaceae). palmata, A. sp., A. flavovaginata); others grow in Opera Bot 126: 1-106. high altitudes (three species of Sphaeradenia, up to FAHN A AND CUTLER DF. 1992. Xerophytes. Encyclopedia 3200 high). These factors could be associated with of plant . Braun H, Carlquist S, Ozenda P and Roth I (Eds), Gebrüder Borntraeger. the anatomical characteristics and their variations, GARCÍA C AND POLAINA J. 2010. Marco conceptual para depending on the species and the place where it productos no maderables del bosque en manglares de grows, so that these anatomical attributes that have Colombia. Gest Ambien 10: 169-178. GUTIÉRREZ IC ET AL. 2007. Prospercción y evaluación de to do with the ability to adaptation and survival especies con potencial de follaje para la elaboración de of the plant and to its ability for a long vase life bouquets. Agron Colom 25: 176-188. (Tuberquia 2007). HARLING GW. 1958. Monograph of the Cyclanthaceae. Acta Horti Berg 18: 1-428. This study provides elements to suggest that HENNY RJ ET AL. 2008. Tropical Foliage Plant Development: the evaluation of different anatomical features of a Breeding Techniques for Anthurium and Spathiphyllum. plant may be useful in the selection of species with Series of Environmental Horticulture Department, p. 1-4. HLWATIKA CN AND BHAT R. 2002. An Ecological potential for use as ornamental foliage. This is just Interpretation of the Difference in Leaf Anatomy and its an approximation to an almost unexplored field of Plasticity in Contrasting Tree Species in Orange Kloof, research with large possibilities in the generation Table Mountain, South Africa. Ann Bot 89: 109-114. KEATING RC. 2014. Preparing plant organs for Light of useful knowledge on sustainable use of plant Microscopic study: A compendium of simple techniques. resources. Missouri Botanical Garden Press.

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